Table Of ContentOxygenic Photosynthesis: The Light Reactions
Advances in Photosynthesis
VOLUME 4
Series Editor:
GOVINDJEE
Department of Plant Biology
University of Illinois, Urbana, Illinois, U.S.A.
Consulting Editors:
Jan AMESZ, Leiden, The Netherlands
Eva-Mari ARO, Turku, Finland
James BARBER, London, United Kingdom
Robert E. BLANKENSHIP, Tempe, Arizona, U.S.A.
Norio MURATA, Okazaki, Japan
Donald R. ORT, Urbana, Illinois, U.S.A.
Advances in Photosynthesis is an ambitious new book series seeking to
provide a comprehensive and state-of-the-art account of photosynthesis
research. Photosynthesis is the process by which higher plants, algae and
certain species of bacteria transform and store solar energy in the form of
energy-rich organic molecules. These compounds are in turn used as the
energy source for all growth and reproduction in these organisms. As such,
virtually all life on the planet ultimately depends on photosynthetic energy
conversion. This series of multiauthored books spans topics from physics to
agronomy, from femtosecond reactions to season long production, from the
photophysics of reaction centers to the physiology of whole organisms, and
from X-ray crystallography of proteins to the morphology of intact plants. The
intent of this new series of publications is to offer beginning researchers,
graduate students, and even research specialists a comprehensive current
picture of the remarkable advances across the full scope of photosynthesis
research.
The titles to be published in this series are listed on the backcover of this volume.
Oxygenic Photosynthesis:
The Light Reactions
Edited by
Donald R. Ort
USDA/ARS, Photosynthesis Research Unit,
University of Illinois,
Urbana, Illinois, U.S.A.
and
Charles F. Yocum
Department of Biology/Chemistry,
University of Michigan,
Ann Arbor, Michigan, U.S.A.
Assistant Editor
Iris F. Heichel
University of Illinois
KLUWER ACADEMIC PUBLISHERS
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eBookISBN: 0-306-48127-8
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Contents
Preface xv
Introduction
1 Electron Transfer and Energy Transduction in Photosynthesis:
An Overview 1–9
D. R. Ort and C. F. Yocum
Summary 1
I. Introduction 2
II. Thylakoid Membranes: Energy Trapping and Electron Transfer Reactions 2
III. Energy Conservation In Thylakoid Membranes 6
IV. Molecular Biology of Thylakoids 7
Acknowledgments 9
References 9
Thylakoid Membranes: Structure
2 Structure, Composition, Functional Organization and Dynamic
Properties of Thylakoid Membranes 11–30
L. Andrew Staehelin and Georg W. M. van der Staay
Summary 11
I. Structural Organization and Compartmentalization of Chloroplasts 12
II. Composition and Distribution of AcylLipids in Thylakoid Membranes 13
III. The Photosynthetic Electron Transport Chain: Components and
Functional Organization 16
IV. Composition of the Five Complexes of the Thylakoid Membrane 18
V. Lateral Distribution of Protein Complexes Between Grana and Stroma
Membrane Regions 18
VI. Macromolecular Organization of Thylakoid Membranes: Correlative
Freeze-fracture, Freeze-etch and Biochemical Studies 19
VII. Why do Thylakoid Membranes Form Grana Stacks? 25
VIII. Alteration in Thylakoid Membrane Composition and Organization in
Response to Changes in the Light Environment 25
Acknowledgments 27
References 27
3 Evolution of Thylakoid Structure 31–40
Gregory R. Wolfe and J. Kenneth Hoober
Summary 31
I. Introduction 31
II. Photosystems 31
III. Light-Harvesting Complexes 35
IV. ThylakoidMorphology 36
V. Conclusions 37
References 38
Thylakoid Membranes: Synthesis and Assembly of Thylakoid
Membranes
4 Control ofThylakoid Membrane Development and Assembly 41–58
AndrewN.Webberand Neil R.Baker
Summary 41
I. Introduction 42
II. Multimeric Protein Complexes 42
III. PhotochemicalActivities 50
IV. Environmental Influences 52
V. Concluding Remarks 53
Acknowledgments 53
References 53
5 Development of Thylakoid Membrane Stacking 59–68
LászlóMustárdy
Summary 59
I. Introduction 60
II. ReconstitutionofThree-dimensional ThylakoidArrangement 60
III. OntogeneticAssembly ofThylakoidsViewedin Three-dimensions 63
IV. PossibleFunctionalRoleofHelicalArrangement 66
References 67
6 Biosynthesis ofThylakoid Membrane Lipids 69–101
Roland DouceandJacquesJoyard
Summary 69
I. Introduction 70
II. Characterization and Biosynthesis ofChloroplastGlycerolipids 70
III. Characterization and Biosynthesis ofChloroplast Prenyllipids 82
IV. Transfer of LipidConstituentsbetween Envelopeand Thylakoids 92
V. Future Prospects 94
Acknowledgments 95
References 95
7 Targeting of Proteins Into and Across theThylakoid Membrane 103–112
Colin Robinson
Summary 103
I. Introduction 104
II. SitesofSynthesisofThylakoid Proteins 104
III. Insertionof Proteinsinto theThylakoid Membrane 104
IV. Translocation ofLumenal ProteinsAcrosstheThylakoid Membrane 107
V. Concluding Remarks 110
References 111
vi
The Photosynthetic Apparatus: Oxygen Evolution
8 Introduction to Oxygen Evolution and the Oxygen-Evolving
Complex 113–136
Terry M. Bricker and Demetrios F. Ghanotakis
Summary 113
I. Introduction 114
II. Photosystem II Preparations 115
III. The Proteins of Photosystem II 116
III. Inorganic Cofactors of Photosystem II 125
IV. The Mechanism of WaterOxidation 128
V. Conclusions 129
Acknowledgments 129
References 129
9 Oxygen Evolution 137–164
R. David Britt
Summary 137
I. Introduction and Overview 138
II. Kinetics and Thermodynamics of Oxygen Evolution 139
III. Structural Studies 141
IV. Manganese Cluster Ligation 148
V. Water Oxidation Mechanisms 154
Acknowledgments 159
References 159
10 Protons and Charge Indicators in Oxygen Evolution 165–192
Michael Haumann and Wolfgang Junge
Summary 165
I. Introduction 166
II. Materials and Techniques 167
III. Extents and Rates of Proton Release During the Redox Transitions 174
IV. Structural and Mechanistic Considerations 184
Acknowledgments 187
References 187
The Photosynthetic Apparatus: Photosystem II
11 Introduction tothe Photosystem II Reaction Center—Isolation and
Biochemical and Biophysical Characterization 193–211
Kimiyuki Satoh
Summary 193
I. Introduction 194
II. Isolation 195
III. Chemical Composition and Polypeptide Sequence 196
IV. Functional Aspects of PS II RC Organization—Energy and Electron
TransferProcesses 200
vii
V. Structural Aspects of Organization—Structure of the Primary Donor (P680) 203
VI. Perspectives 207
References 207
12 Structure, Dynamics, and Energy Conversion Efficiency in
Photosystem II 213–247
Bruce A. Diner and Gerald T. Babcock
Summary 214
I. Introduction 214
II. Biochemical and Pigment Organization of Photosystem II 218
III. Absorption Characteristics and Organization of Pigments in Photosystem II 218
IV. Primary Photochemistry 222
V. Secondary Electron Transfer 226
Acknowledgments 240
References 240
13 Form and Function of Cytochrome b-559 249–264
John Whitmarsh and Himadri B. Pakrasi
Summary 249
I. Introduction 250
II. Cytochrome b-559 Structure 250
III. Cytochrome b-559 Forms in Photosystem II 252
IV. Cytochrome b-559 Function 257
Acknowledgments 260
References 260
14 Photosystem II Heterogeneity 265–287
Jérôme Lavergne and Jean-Marie Briantais
Summary 265
I. Introduction 266
II. Static Heterogeneities: Basic Data 267
III. Possible Correlations 274
IV. Dynamic Heterogeneities 276
VI. Conclusions 281
Acknowledgments 281
References 282
The Photosynthetic Apparatus: Photosystem I
15 Introduction to Photosystem I: Reaction Center Function,
Composition and Structure 289–311
Rachel Nechushtai, Amir Eden, Yuval Cohen and Judith Klein
Summary 290
I. Introduction 290
II. The Composition and Function of PS I Components 292
III. The Organization and Structure of Photosystem I 298
Acknowledgments 306
References 306
viii
16 Photosystem I Electron Transfer Reactions—Components and
Kinetics 313–332
Richard Malkin
Summary 313
I. Introduction 314
II. Electron Carriers in PS I 314
III. Kinetics of PS I Electron Transfer Reactions 321
IV. Cyclic Electron Transfer and Cyclic Phosphorylation 327
Acknowledgments 329
References 329
17 Ferredoxin and Ferredoxin-Dependent Enzymes 333–361
David B. Knaff
Summary 333
I. Introduction 334
II. Ferredoxin 334
III. Ferredoxin:NADP+ Oxidoreductase (FNR) 339
IV. Nitrite Reductase 346
V. Glutamate Synthase 349
VI. Thioredoxin Reductase 351
VII. Conclusion 353
Acknowledgments 353
References 354
18 Structure Analysis of Single Crystals of Photosystem I
by X-Ray, EPR and ENDOR: A Short Status Report 363–375
H. T. Witt
Summary 363
I. Introduction 364
II. The PS I Complex of Cyanobacterium Synechococcuselongatus 364
III. Crystallization of the Trimer of PS I 365
IV. X-Ray Structure Analysis of Single Crystals of PS I at 4.5 Å Resolution 365
V. EPR Structure Analysis of P700 and the Fe-S clusters in PS I Complexes
and Single Crystals at Low Temperature 370
VI. ENDOR Structure Analysis of P700 in Single Crystals of PS I at
Low Temperature 371
Acknowledgments 374
References 374
The Photosynthetic Apparatus: Components of Intersystem Elec-
tronTransfer
19 The Cytochrome Complex—Composition, Structure
and Function 377–398
Günter Hauska, Michael Schütz and Michael Büttner
Summary 377
I. Introduction 378
II. Occurrence 378
III. Isolation 380
ix
IV. Components and Structure 380
V. Function 387
VI. Regulation 391
VIII. Conclusions and Open Questions 392
Acknowledgments 393
Note in Proof 394
References 394
20 Basic Aspects of Electron and Proton Transfer Reactions with
Applications to Photosynthesis 399–411
Lev I. Krishtalik and William A. Cramer
Summary 399
I. Introduction 400
II. Medium Reorganization in Electron Transfer 400
III. Transition Probability 402
IV. Proteins As Polar Media 402
V. Proteins As an Intervening Medium in Electron Transfer 403
VI. Pathways of Proton Transfer in Energy-Transducing Membranes 407
Acknowledgments 409
References 409
21 Plastocyanin: Structure, Location, Diffusion and Electron
Transfer Mechanisms 413–429
Elizabeth L. Gross
Summary 413
I. Introduction 414
II. Structure 414
III. The Interaction of Plastocyanin with Cytochrome f 420
IV. The Interaction of Plastocyanin with P700 423
V. The Function of Plastocyanin in the Lumen of the Thylakoid at Acid pH 425
VI. Conclusions 426
Acknowledgments 426
References 426
22 Some Consequences of the High Resolution X-Ray Structure
Analysis of Cytochrome f 431–437
S. E. Martinez, D. Huang, J. L. Smith and W. A. Cramer
Summary 431
I. Introduction 431
II. Preparation and Crystallization of TruncatedCytochrome f 432
III. Unprecedented Aspects of the Structure 434
IV. Other Major Properties of the Structure 434
V. Orientation of Cytochrome f Relative to the Membrane Plane 436
VI. Evolutionary Relation between Cytochromes f and 436
VII. Electron Transferfrom Cyt f to Plastocyanin 436
Acknowledgments 437
References 437
Summary 439
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